Slime mold or slime mould is an informal name given to a polyphyletic assemblage of unrelated eukaryotic organisms in the Stramenopiles, Rhizaria, Discoba, Amoebozoa and Holomycota clades. Most are microscopic; those in the Myxogastria form larger plasmodial slime molds visible to the naked eye. The slime mold life cycle includes a free-living single-celled stage and the formation of spores. Spores are often produced in macroscopic multicellular or multinucleate fruiting bodies that may be formed through aggregation or fusion; aggregation is driven by chemical signals called acrasins. Slime molds contribute to the decomposition of dead vegetation; some are parasitic.
Most slime molds are terrestrial and free-living, typically in damp shady habitats such as in or on the surface of rotting wood. Some myxogastrians and protostelians are aquatic or semi-aquatic. The phytomyxea are parasitic, living inside their plant hosts. Geographically, slime molds are cosmopolitan in distribution. A small number of species occur in regions as dry as the Atacama Desert and as cold as the Arctic; they are abundant in the tropics, especially in rainforests.
Slime molds have a variety of behaviors otherwise seen in animals with brains. Species such as Physarum polycephalum have been used to simulate traffic networks. Some species have traditionally been eaten in countries such as Ecuador.
The first account of slime molds was Thomas Panckow 's 1654 discussion of Lycogala epidendrum . He called it Fungus cito crescentes, "a fast-growing fungus". [2] [1]
German mycologist Heinrich Anton de Bary, in 1860 and 1887, classified the Myxomycetes (plasmodial slime molds) and Acrasieae (cellular slime molds) as Mycetozoa, a new class. He also introduced a "Doubtful Mycetozoa" section for Plasmodiophora (now in Phytomyxea) and Labyrinthula , emphasizing their distinction from plants and fungi. [3] [4] In 1880, the French botanist Philippe van Tieghem analyzed the two groups further. [4] In 1868, the German biologist Ernst Haeckel placed the Mycetozoa in a kingdom he named Protista. [4] In 1885, the British zoologist Ray Lankester grouped the Mycetozoa alongside the Proteomyxa as part of the Gymnomyxa in the phylum Protozoa. [4] Arthur and Gulielma Lister published monographs of the group in 1894, 1911, and 1925. [5] [6]
In 1932 and 1960, the American mycologist George Willard Martin argued that the slime molds evolved from fungi. [7] [8] In 1956, the American biologist Herbert Copeland placed the Mycetozoa (the myxomycetes and plasmodiophorids) and the Sarkodina (the labyrinthulids and the cellular slime molds) in a phylum called Protoplasta, which he placed alongside the fungi and the algae in a new kingdom, Protoctista. [4] [9]
In 1969, the taxonomist R. H. Whittaker observed that slime molds were highly conspicuous and distinct within the Fungi, the group to which they were then classified. He concurred with Lindsay S. Olive's proposal to reclassify the Gymnomycota, which includes slime molds, as part of the Protista. [10] Whittaker placed three phyla, namely the Myxomycota, Acrasiomycota, and Labyrinthulomycota in a subkingdom Gymnomycota within the Fungi. [4] The same year, Martin and Alexopoulos published their influential textbook The Myxomycetes. [6]
In 1975, Olive distinguished the dictyostelids and the acrasids as separate groups. [4] In 1992, David J. Patterson and M. L. Sogin proposed that the dictyostelids diverged before plants, animals, and fungi. [11]
Slime molds have little or no fossil history, as might be expected given that they are small and soft-bodied. [12] The grouping is polyphyletic, consisting of multiple clades (emphasised in the phylogenetic tree) widely scattered across the Eukaryotes. Paraphyletic groups are shown in quotation marks: [13] [14]
Various estimates of the number of species of slime molds agree that there are around 1000 species, most being Myxogastria. Collection of environmental DNA gives a higher estimate, from 1200 to 1500 species. [6] These are diverse both taxonomically and in appearance, the largest and most familiar species being among the Myxogastria. The growth forms most commonly noticed are the sporangia, the spore-forming bodies, which are often roughly spherical; these may be directly on the surface, such as on rotting wood, or may be on a thin stalk which elevates the spores for release above the surface. Other species have the spores in a large mass, which may be visited by insects for food; they disperse spores when they leave. [15]
The Myxogastria or plasmodial slime molds are the only macroscopic scale slime molds; they gave the group its informal name, since for part of their life cycle they are slimy to the touch. [16] A myxogastrian consists of a large cell with thousands of nuclei within a single membrane without walls, forming a syncytium. [17] Most are smaller than a few centimeters, but some species may reach sizes up to several square meters, and in the case of Brefeldia maxima , a mass of up to 20 kilograms (44 lb). [18] [19] [20]
The Dictyosteliida or cellular slime molds do not form huge coenocytes like the Myxogastria; their amoebae remain individual for most of their lives as individual unicellular protists, feeding on microorganisms. When food is depleted and they are ready to form sporangia, they form swarms. The amoebae join up into a tiny multicellular slug which crawls to an open lit place and grows into a fruiting body, a sorocarp. Some of the amoebae become spores to begin the next generation, but others sacrifice themselves to become a dead stalk, lifting the spores up into the air. [23] [24]
The Protosteliida, a polyphyletic group, have characters intermediate between the previous two groups, but they are much smaller, the fruiting bodies only forming one to a few spores. [25]
The lobosans, a paraphyletic group of amoebae, include the Copromyxa slime molds. [26] [27]
Among the non-amoebozoan slime molds are the Acrasids, which have sluglike amoebae. In locomotion, the amoebae's pseudopodia are eruptive, meaning that hemispherical bulges appear at the front. [28] The Phytomyxea are obligate parasites, with hosts among the plants, diatoms, oomycetes, and brown algae. They cause plant diseases like cabbage club root and powdery scab. [29] The Labyrinthulomycetes are marine slime nets, forming labyrinthine networks of tubes in which amoeba without pseudopods can travel. [30] The Fonticulida are cellular slime molds that form a fruiting body in a "volcano" shape. [31]
Slime molds, with their small size and moist surface, live mostly in damp habitats including shaded forests, rotting wood, fallen or living leaves, and on bryophytes. [32] [18] Most Myxogastria are terrestrial, [18] though some, like Didymium aquatilis are aquatic, [33] [34] and D. nigripes is semi-aquatic. [34] Myxogastria are not limited to wet regions; 34 species are known from Saudi Arabia, living on bark, in plant litter, and rotting wood, even in deserts. [35] They occur, too, in Arizona's Sonoran Desert (46 species), and in Chile's exceptionally dry Atacama Desert (24 species). In contrast, the semi-dry Tehuacán-Cuicatlán Biosphere Reserve has 105 species, and Russia and Kazakhstan's Volga river basin has 158 species. [35] In tropical rainforests of Latin America, species such as of Arcyria and Didymium are commonly epiphyllous, growing on the leaves of liverworts. [36]
The dictyostelids are mostly terrestrial. [37] On Changbai Mountain in China, six species of dictyostelids were found in forest soils at elevations up to 2,038 m (6,686 ft), the highest recorded species there being Dictyostelium mucoroides. [38] The protostelids live mainly on dead plant matter, where they consume the spores of bacteria, yeasts, and fungi. [37] They include some aquatic species, which live on dead plant parts submerged in ponds. [33] Cellular slime molds are most numerous in the tropics, decreasing with latitude, but are cosmopolitan in distribution, occurring in soil even in the Arctic and the Antarctic. [39] In the Alaskan tundra, the only slime molds are the dictyostelids D. mucoroides and D. sphaerocephalum. [36]
The species of Copromyxa are coprophilous, feeding on dung. [27]
Some myxogastrians have their spores dispersed by animals. The slime mold fly Epicypta testata lay its eggs within the spore mass of Enteridium lycoperdon , which the larvae feed on. These pupate, and the hatching adults carry and disperse spores that have stuck to them. [21] While various insects consume slime molds, Sphindidae slime mold beetles, both larvae and adults, exclusively feed on them. [40]
Plasmodial slime molds begin life as amoeba-like cells. These unicellular amoebae are commonly haploid and feed on small prey such as bacteria, yeast cells, and fungal spores by phagocytosis, engulfing them with its cell membrane. These amoebae can mate if they encounter the correct mating type and form zygotes that then grow into plasmodia. These contain many nuclei without cell membranes between them, and can grow to meters in size. The species Fuligo septica is often seen as a slimy yellow network in and on rotting logs. The amoebae and the plasmodia engulf microorganisms. [41] The plasmodium grows into an interconnected network of protoplasmic strands. [42] Within each protoplasmic strand, the cytoplasmic contents rapidly stream, periodically reversing direction. The streaming protoplasm within a plasmodial strand can reach speeds of up to 1.35 mm per second in Physarum polycephalum, the fastest for any microorganism. [43]
Slime molds are isogamous, which means that their gametes (reproductive cells) are all the same size, unlike the eggs and sperms of animals. [45] Physarum polycephalum has three genes involved in reproduction: matA and matB, with thirteen variants each, and matC with three variants. Each reproductively mature slime mold is diploid, meaning that it contains two copies of each of the three reproductive genes. [46] When P. polycephalum is ready to make its reproductive cells, it grows a bulbous extension of its body to contain them. [47] Each cell has a random combination of the genes that the slime mold contains within its genome. Therefore, it can create cells of up to eight different gene types. Released cells then independently seek another compatible cell for fusion. Other individuals of P. polycephalum may contain different combinations of the matA, matB, and matC genes, allowing over 500 possible variations. It is advantageous for organisms with this type of reproductive cell to have many mating types because the likelihood of the cells finding a partner is greatly increased, and the risk of inbreeding is drastically reduced. [46]
The cellular slime molds are a group of approximately 150 described species. They occur primarily in the humus layer of forest soils [48] and feed on bacteria but also are found in animal dung and agricultural fields. They exist as single-celled organisms while food is plentiful. When food is in short supply, many of the single-celled amoebae congregate and start moving as a single body, called a 'slug'. The ability of the single celled organisms to aggregate into multicellular forms are why they are also called the social amoebae. In this state they are sensitive to airborne chemicals and can detect food sources. They readily change the shape and function of parts, and may form stalks that produce fruiting bodies, releasing countless spores, light enough to be carried on the wind or on passing animals. [23] The cellular slime mold Dictyostelium discoideum has many different mating types. When this organism has entered the stage of reproduction, it releases a chemical attractant. [49] When it comes time for the cells to fuse, Dictyostelium discoideum has mating types of its own that dictate which cells are compatible with each other. There are at least eleven mating types; macrocysts form after cell contact between compatible mating types. [50]
The chemicals that aggregate cellular slime molds are small molecules called acrasins; motion towards a chemical signal is called chemotaxis. The first acrasin to be discovered was cyclic adenosine monophosphate (cyclic AMP), a common cell signaling molecule, in Dictyostelium discoideum. During the aggregation phase of their life cycle, Dictyostelium discoideum amoebae communicate with each other using traveling waves of cyclic AMP. [51] [52] [53] There is an amplification of cyclic AMP when they aggregate. [54] Pre-stalk cells move toward cyclic AMP, but pre-spore cells ignore the signal. [55] Other acrasins exist; the acrasin for Polysphondylium violaceum , purified in 1983, is the dipeptide glorin. [56] Calcium ions too serve to attract slime mold amoebae, at least at short distances. It has been suggested that acrasins may be taxon-specific, since specificity is required to form an aggregation of genetically similar cells. Many dictyostelid species indeed do not respond to cyclic AMP, but as of 2023 their acrasins remained unknown. [57]
The practical study of slime molds was facilitated by the introduction of the "moist culture chamber" by H. C. Gilbert and G. W. Martin in 1933. [58] Slime molds can be used to teach convergent evolution, as the habit of forming a stalk with a sporangium that can release spores into the air, off the ground, has evolved repeatedly, such as in myxogastria (eukaryotes) and in myxobacteria (prokaryotes). [59] Further, both the (macroscopic) dictyostelids and the (microscopic) protostelids have a phase with motile amoebae and a phase with a stalk; in the protostelids, the stalk is tiny, supporting just one spore, but the logic of airborne spore dispersal is the same. [59]
O. R. Collins showed that the slime mold Didymium iridis had two strains (+ and −) of cells, equivalent to gametes, that these could form immortal cell lines in culture, and that the system was controlled by alleles of a single gene. This made the species a model organism for exploring incompatibility, asexual reproduction, and mating types. [59]
Slime molds have been studied for their production of unusual organic compounds, including pigments, antibiotics, and anti-cancer drugs. [59] Pigments include naphthoquinones, physarochrome A, and compounds of tetramic acid. Bisindolylmaleimides produced by Arcyria denudata include some phosphorescent compounds. [60] The sporophores (fruiting bodies) of Arcyria denudata are colored red by arcyriaflavins A–C, which contain an unusual indolo[2,3-a]carbazole alkaloid ring. [61] By 2022, more than 100 pigments had been isolated from slime molds, mostly from sporophores. It has been suggested that the many yellow-to-red pigments might be useful in cosmetics. [15] Some 42% of patients with seasonal allergic rhinitis reacted to myxogastrian spores, so the spores may contribute significantly as airborne allergens. [62]
Slime molds share some similarities with neural systems in animals. [63] The membranes of both slime molds and neural cells contain receptor sites, which alter electrical properties of the membrane when it is bound. [64] Therefore, some studies on the early evolution of animal neural systems are inspired by slime molds. [65] [66] [67] When a slime mold mass or mound is physically separated, the cells find their way back to re-unite. Studies on Physarum polycephalum have even shown the organism to have an ability to learn and predict periodic unfavorable conditions in laboratory experiments. [68] John Tyler Bonner, a professor of ecology known for his studies of slime molds, argues that they are "no more than a bag of amoebae encased in a thin slime sheath, yet they manage to have various behaviors that are equal to those of animals who possess muscles and nerves with ganglia – that is, simple brains." [69]
The slime mold algorithm is a meta-heuristic algorithm, based on the behavior of aggregated slime molds as they stream in search of food. It is described as a simple, efficient, and flexible way of solving optimization problems, such as finding the shortest path between nodes in a network. However, it can become trapped in a local optimum. [70]
Toshiyuki Nakagaki and colleagues studies slime molds and their abilities to solve mazes by placing nodes at two point separated by a maze of plastic film. The mold explored all possible paths and solves it for the shortest path. [71]
Atsushi Tero and colleagues grew Physarum in a flat wet dish, placing the mold in a central position representing Tokyo, and oat flakes surrounding it corresponding to the locations of other major cities in the Greater Tokyo Area. As Physarum avoids bright light, light was used to simulate mountains, water and other obstacles in the dish. The mold first densely filled the space with plasmodia, and then thinned the network to focus on efficiently connected branches. The network closely resembled Tokyo's rail system. [72] [73] P. polycephalum was used in experimental laboratory approximations of motorway networks of 14 geographical areas: Australia, Africa, Belgium, Brazil, Canada, China, Germany, Iberia, Italy, Malaysia, Mexico, the Netherlands, UK and US. [74] [75] [76] The filamentary structure of P. polycephalum forming a network to food sources is similar to the large scale galaxy filament structure of the universe. This observation has led astronomers to use simulations based on the behaviour of slime molds to inform their search for dark matter. [77] [78]
In central Mexico, the false puffball Enteridium lycoperdon was traditionally used as food; it was one of the species which mushroom-collectors or hongueros gathered on trips into the forest in the rainy season. One of its local names is "cheese mushroom", so called for its texture and flavor when cooked. It was salted, wrapped in a maize leaf, and baked in the ashes of a campfire; or boiled and eaten with maize tortillas. Fuligo septica was similarly collected in Mexico, cooked with onions and peppers and eaten in a tortilla. In Ecuador, Lycogala epidendrum was called "yakich" and eaten raw as an appetizer. [79]
Oscar Requejo and N. Floro Andres-Rodriguez suggest that Fuligo septica may have inspired Irvin Yeaworth's 1958 film The Blob , in which a giant amoeba from space sets about engulfing people in a small American town. [79]
The dictyostelids or cellular slime molds are a group of slime molds or social amoebae.
Mycetozoa is a polyphyletic grouping of slime molds. It was originally thought to be a monophyletic clade, but recently it was discovered that protostelia are a polyphyletic group within Conosa.
Dictyostelium is a genus of single- and multi-celled eukaryotic, phagotrophic bacterivores. Though they are Protista and in no way fungal, they traditionally are known as "slime molds". They are present in most terrestrial ecosystems as a normal and often abundant component of the soil microflora, and play an important role in the maintenance of balanced bacterial populations in soils.
Amoebozoa is a major taxonomic group containing about 2,400 described species of amoeboid protists, often possessing blunt, fingerlike, lobose pseudopods and tubular mitochondrial cristae. In traditional classification schemes, Amoebozoa is usually ranked as a phylum within either the kingdom Protista or the kingdom Protozoa. In the classification favored by the International Society of Protistologists, it is retained as an unranked "supergroup" within Eukaryota. Molecular genetic analysis supports Amoebozoa as a monophyletic clade. Modern studies of eukaryotic phylogenetic trees identify it as the sister group to Opisthokonta, another major clade which contains both fungi and animals as well as several other clades comprising some 300 species of unicellular eukaryotes. Amoebozoa and Opisthokonta are sometimes grouped together in a high-level taxon, named Amorphea. Amoebozoa includes many of the best-known amoeboid organisms, such as Chaos, Entamoeba, Pelomyxa and the genus Amoeba itself. Species of Amoebozoa may be either shelled (testate) or naked, and cells may possess flagella. Free-living species are common in both salt and freshwater as well as soil, moss and leaf litter. Some live as parasites or symbionts of other organisms, and some are known to cause disease in humans and other organisms.
Physarum polycephalum, an acellular slime mold or myxomycete popularly known as "the blob", is a protist with diverse cellular forms and broad geographic distribution. The “acellular” moniker derives from the plasmodial stage of the life cycle: the plasmodium is a bright yellow macroscopic multinucleate coenocyte shaped in a network of interlaced tubes. This stage of the life cycle, along with its preference for damp shady habitats, likely contributed to the original mischaracterization of the organism as a fungus. P. polycephalum is used as a model organism for research into motility, cellular differentiation, chemotaxis, cellular compatibility, and the cell cycle.
The family Acrasidae is a family of slime molds which belongs to the excavate group Percolozoa. The name element acrasio- comes from the Greek akrasia, meaning "acting against one's judgement". This group consists of cellular slime molds.
Eumycetozoa, or true slime molds, is a diverse group of protists that behave as slime molds and develop fruiting bodies, either as sorocarps or as sporocarps. It is a monophyletic group or clade within the phylum Amoebozoa that contains the myxogastrids, dictyostelids and protosporangiids.
Each species of slime mold has its own specific chemical messenger, which are collectively referred to as acrasins. These chemicals signal that many individual cells aggregate to form a single large cell or plasmodium. One of the earliest acrasins to be identified was cyclic AMP, found in the species Dictyostelium discoideum by Brian Shaffer, which exhibits a complex swirling-pulsating spiral pattern when forming a pseudoplasmodium.
Multinucleate cells are eukaryotic cells that have more than one nucleus, i.e., multiple nuclei share one common cytoplasm. Mitosis in multinucleate cells can occur either in a coordinated, synchronous manner where all nuclei divide simultaneously or asynchronously where individual nuclei divide independently in time and space. Certain organisms may have a multinuclear stage of their life cycle. For example, slime molds have a vegetative, multinucleate life stage called a plasmodium.
Dictyostelium discoideum is a species of soil-dwelling amoeba belonging to the phylum Amoebozoa, infraphylum Mycetozoa. Commonly referred to as slime mold, D. discoideum is a eukaryote that transitions from a collection of unicellular amoebae into a multicellular slug and then into a fruiting body within its lifetime. Its unique asexual life cycle consists of four stages: vegetative, aggregation, migration, and culmination. The life cycle of D. discoideum is relatively short, which allows for timely viewing of all stages. The cells involved in the life cycle undergo movement, chemical signaling, and development, which are applicable to human cancer research. The simplicity of its life cycle makes D. discoideum a valuable model organism to study genetic, cellular, and biochemical processes in other organisms.
Protosteliomycetes/Protosteliales (ICBN) or Protostelea/Protostelia/Protosteliida (ICZN) is a grouping of slime molds from the phylum Mycetozoa. The name can vary depending upon the taxon used. Other names include Protostelea, Protostelia, and Protostelida. When not implying a specific level of classification, the term protostelid or protosteloid amoeba is sometimes used.
Myxogastria/Myxogastrea or Myxomycetes (ICN) is a class of slime molds that contains 5 orders, 14 families, 62 genera, and 888 species. They are colloquially known as the plasmodial or acellular slime moulds.
Differentiation-inducing factor (DIF) is one of a class of effector molecules that induce changes in cell chemistry, inhibiting growth and promoting differentiation of cell type. This name has been given to several factors before it was clear if they were the same or different effectors. DIFs have garnered interest with their potential tumor inhibiting properties. DIFs have also been used to help regulate plant growth.
Trichiales is an order of slime moulds in the phylum Amoebozoa. Trichiales is one of five orders in the group Myxomycetes, or the true plasmodial slime molds. It is also currently categorized under the superorder Lucisporidia with its sister group, Liceales. The order was first described by Thomas MacBride in 1922, and has retained the same name and status as a defined order in present phylogeny. In the plasmodium form, members of Trichiales lack a columella but have a well-developed capillitium for spore dispersal. The shape and details of the capillitium are used to define families within the order. Spores are brightly coloured, ranging from clear, white and yellow to pink and red-brown tones. The order currently has 4 families, 14 genera and 174 species. Recent molecular research has shown that while Trichiales probably represents a true taxonomic group, its sister group Liceales is likely paraphyletic, and it has been suggested that several genera from the Liceales should be reclassified under Trichiales instead.
Fonticula is a genus of cellular slime mold which forms a fruiting body in a volcano shape. As long ago as 1979 it has been known to not have a close relationship with either the Dictyosteliida or the Acrasidae, the two well-established groups of cellular slime molds. In 1979, Fonticula was made a new genus of its own due to the unique characteristics of its fruiting body, with only one species: Fonticula alba.
Enteridium lycoperdon, the false puffball, is one of the more obvious species of slime mould or Myxogastria, typically seen in its reproductive phase as a white 'swelling' on standing dead trees in the spring, or on large pieces of fallen wood. Alder is a common host.
Polysphondylium pallidum is a species of cellular slime mould, a member of the phylum Mycetozoa.
In true slime molds (myxogastria), lichens, and in species of the family Clavicipitaceae, the hypothallus is the layer on which the fruit body sits, lying in contact with the substrate. The word is derived from the Ancient Greek root hypó ("under") and thallós.
An amoeba, often called an amoeboid, is a type of cell or unicellular organism with the ability to alter its shape, primarily by extending and retracting pseudopods. Amoebae do not form a single taxonomic group; instead, they are found in every major lineage of eukaryotic organisms. Amoeboid cells occur not only among the protozoa, but also in fungi, algae, and animals.
Hemitrichia is a genus of slime molds, of the family Trichiidae, found within the order Trichiida. It was first described by Josef Rostafinski in 1873 and remains a well-defined genus of the slime molds. Hemitrichia species exhibit either plasmodiocarp or sporangium fruiting bodies, both of which are well-known and recognizable slime molds seen on multiple continents. The genus includes Hemtrichia serpula, known as the pretzel slime mold, an iconic and widespread species that has been used to examine speciation in slime molds.